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A NONMARINE VERTEBRATE COPROLITE ACME ZONE IN THE PERMO-TRIASSIC

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Lucas, S.G. and Zeigler, K.E., eds., 2005, The Nonmarine Permian, New Mexico Museum of Natural History and Science Bulletin No. 30.
INTRODUCTION
Häntzschel et al. (1968, fig. 3; Fig. 1) published the only statis-
tical analysis of the frequency of coprolites through time. This analysis
included both invertebrate and vertebrate coprolites from all environ-
ments and was based upon “statistical counting of the geologic age
assigned to the coprolite and faecal pellet specimens described in the
literature” (Häntzschel et al., 1968, p. 4). This figure shows a strong
correlation between frequency and age, with younger strata yielding
more coprolites. Häntzschel et al. (1968, p. 4) cautioned that this analysis
depended “widely upon the scientific advancements and activities of
geologists and paleontologists in the various countries of the world”
and “does not necessarily represent the actual distribution of coprolites
in geological time.”
Hunt et al. (1994, p. 2273-224) provided a brief qualitative re-
view of the distribution of vertebrate coprolites through the Phanero-
zoic. They noted, for example, that herbivore coprolites are rare prior
to the Quaternary. The purpose of this paper is to briefly discuss the
obvious abundance of vertebrate coprolites in the Permo-Triassic.
PHANEROZOIC RECORD OF NONMARINE VERTEBRATE
COPROLITES
Over the last 30 years, we have conducted extensive fieldwork
in nonmarine strata in North America, Europe and Asia ranging in age
from Carboniferous to Quaternary. It is this experience in addition to
our knowledge of museum collections that forms the basis of the fol-
lowing generalizations.
We have always been aware that “most paleontologists do not
collect vertebrate coprolites” (Hunt et al., 1994, p. 224). Vertebrate
coprolites are often the most common vertebrate trace fossils in non-
eolian environments (e.g., Hunt et al., 1998). However, this abundance
is not reflected in most paleontological collections, principally because
the majority of vertebrate paleontologists are principally interested in
body fossils (Hunt et al., 1994). Possibly the only major collection in
proportion to the number of vertebrate coprolites encountered by its
field collectors, is the New Mexico Museum of Natural History and
Science.
Our qualitative assessment of the distribution of nonmarine ver-
tebrate coprolites through the Phanerozoic suggests that pre-Permian
coprolites are uncommon. However, coprolites are abundant locally in
Permo-Triassic redbeds (e.g., Hunt and Lucas, 2005a, b). Later Meso-
zoic strata yield few coprolites. Coprolites are locally common in the
Tertiary (references in Hunt et al., 1994). Quaternary coprolites are
abundant, but only in certain depositional settings, notably caves (e.g.,
Buckland, 1823).
PERMO-TRIASSIC ACME FOR NONMARINE
VERTEBRATE COPROLITES
Early Permian-Late Triassic redbeds yield abundant vertebrate
coprolites— they have been reported from most significant outcrops of
strata of this age, e.g., North America (Neumayer, 1904; Hunt et al.,
1998), Europe (Augusta, 1936) and Asia (Jain, 1983; Ochev, 1974).
Vertebrate coprolites are not pervasive through all facies and are
only locally abundant (Hunt et al., 1988). The relative abundance of
different trace fossils is nonrandom and in need of further study. For
example, the Early Permian (principally Wolfcampian) redbeds of New
Mexico yield abundant tetrapod, and lesser numbers of invertebrate,
tracks and yet vertebrate coprolites are relatively uncommon. How-
ever, the reverse is true in the Early Permian (Wolfcampian-Leonardian)
redbeds of Texas. Notably, there are very large collections of vertebrate
trace fossils from both New Mexico and Texas, so sample bias is not a
significant factor in this disparity. Preliminary study of Late Triassic
coprolite distribution indicates that distinct coprofacies can be discrimi-
nated in the Chinle Group of western North America (Hunt et al., 1998).
A NONMARINE VERTEBRATE COPROLITE ACME ZONE IN THE PERMO-TRIASSIC
ADRIAN P. HUNT AND SPENCER G. LUCAS
New Mexico Museum of Natural History, 1801 Mountain Road NW, Albuquerque, NM 87104-1375
Abstract—Nonmarine vertebrate coprolites are uncommon in the pre-Permian, locally abundant in Permo-
Triassic redbeds, rare in the later Mesozoic strata, locally common in the Tertiary and common in certain
depositional settings (e. g., caves) in the Quaternary. There is thus a Permo-Triassic acme zone for vertebrate
coprolites in nonmarine environments.
FIGURE 1. Assessment of the distribution of coprolites through time by Häntzschel
et al. (1968, fig. 3).
Augusta, J., 1936, Ein Stegocephalian-Koprolith aus dem mährischen Perm:
Zentralblatt für Mineralogie, Geologie und Pälaontologie, Abheinlung B, 1936,
p. 334-337.
Buckland, W., 1823, Reliquiae diluvianae: London, 303 p.
Häntzschel, W., El-Baz, F. and Amstutz, G. C., 1968, Coprolites: An annotated
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Hunt, A. P. and Lucas, S. G., 2005a, A new coprolite ichnotaxon from the Early
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Hunt, A. P., Chin, K. and Lockley, M. G., 1994, The paleobiology of coprolites; in
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Jain, S. L., 1983, Spirally-coiled coprolites from the Upper Triassic Maleri Forma-
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51, p. 121-128.
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logical Journal, v. 194, p. 253-255.
... They noted occurrences at 19 NPS units, and we are now aware of an additional 10 records. Other papers in this volume review aspects of the Arizona trace fossil record by time period (Elliott and Blakey, 2005; Heckert et al, 2005; Hunt et al., 2005; Lucas et al., 2005; Lucas and Heckert, 2005; Morgan and White, 2005). USNM refers to the United States National Museum (Smithsonian) in Washington; MNA refers to the Museum of Northern Arizona in Flagstaff; NMMNH refers to the New Mexico Museum of Natural History and Science in Albuquerque. ...
... These specimens pertain to Chelichnus bucklandi and C. duncani and include parallel trackways (Kramer et al., 1995; Lockley and Hunt, 1995, figs. 2.11; Hunt et al., 2005). ...
... The ichnofauna of the eolian DeChelly Sandstone is broadly similar to that of the Coconino Sandstone in being dominated by Chelichnus (C. bucklandi and C. duncani)(Hunt et al., 2005). The DeChelly is unusual among Permian eolianites in yielding a specimen of the lacertoid track Dromopus cf. ...
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Arizona has significant tetrapod ichnofaunas, many of which are from National Park Service units, including traces from the Pennsylvanian Wescogame Formation, Permian Coconino and DeChelly sandstones and Hermit Formation, Triassic Moenkopi Formation and Blue Mesa and Sonsela Members of the Petrified Forest Formation, Jurassic Navajo Sandstone, Cretaceous Toreva Formation, Miocene Bidahochi Formation and the Pliocene Verde Formation. Arizona ichnofaunas are significant for several reasons as they include the first large Paleozoic ichnofaunas described, westernmost Pennsylvanian tetra-pod tracks in North America, largest collected and described sample sizes of trace fossils from eolianites, the most significant Early-Middle Triassic tetrapod ichnofaunas in the New World, and a Cretaceous dinosaur tracksite with multiple tail drags. Other vertebrate trace fossils from Arizona include coprolites from the Moenkopi Formation and Chinle Group and late Cenozoic cave deposits, putative nests from the Chinle Group and numerous middens from the late Pleistocene. There are four temporal phases in the taphonomy of tetrapod tracks: Devonian, Carboniferous-Triassic, Jurassic-Cretaceous and Cenozoic.
... This coprolite abundance is unexpected and particularly significant because of the rarity of pre-Permian coprolites from nonmarine environments (Hunt and Lucas, 2005a), as well as the heretofore unknown occurrence of Pennsylvanian coprolites from a laterally extensive nonmarine facies . Indeed, the high abundance and three-dimensional preservation of coprolites at the Ojo de Amado and Cerrillos del Coyote localities is comparable only to younger vertebrate coprofaunas of the nonmarine Permo-Triassic coprolite acme zone (Hunt and Lucas, 2005a). ...
... This coprolite abundance is unexpected and particularly significant because of the rarity of pre-Permian coprolites from nonmarine environments (Hunt and Lucas, 2005a), as well as the heretofore unknown occurrence of Pennsylvanian coprolites from a laterally extensive nonmarine facies . Indeed, the high abundance and three-dimensional preservation of coprolites at the Ojo de Amado and Cerrillos del Coyote localities is comparable only to younger vertebrate coprofaunas of the nonmarine Permo-Triassic coprolite acme zone (Hunt and Lucas, 2005a). ...
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We document a rare, laterally extensive nonmarine coprofauna preserved in lacustrine sediments of the Upper Pennsylvanian (Missourian) Tinajas Member of the Atrasado Formation, east of Socorro, New Mexico. Abundant coprolites from five recently discovered localities in the Tinajas black shale interval are distinguished from those of the stratigraphically equivalent Tinajas Lagerstätte by their large size and three-dimensional preservation. Together, the Tinajas black shale coprolites represent a remarkable record of morphological and preservational variation within the context of a single paleolake system. Three coprolite taphotypes are recognized: (1) enclosure within a goethite concretionary layer; (2) cementation within a quartz-rich mantle; and (3) non-concretionary hydroxylapatite permineralization. X-ray diffraction analysis (XRD) was used to determine the mineralogical composition of a representative sample of coprolites from each taphotype. Early diagenetic (pre-compaction) mineralization and nucleation of coprolites at the sediment-water interface was a function of possible microbial activity and prevailing geochemical conditions such as iron and silica activity, pH, and oxygen fugacity of lake bottom water and sediment at the time of deposition. Infilling of barite within the cracks of some coprolites represents either microbial sulfate reduction within the microenvironment of the original fecal material or fault-related secondary mineralization that occurred long after deposition.
... Mesozoic vertebrate coprolites are widely distributed in non-marine sediments (e.g., Chin and Gill, 1996;Chin and Kirkland, 1998;Ghosh et al., 2003;Kar et al., 2004;Ambwani and Dutta, 2005;Prasad et al., 2005;Chin, 2007;Dutta and Ambwani, 2007;Chin et al., 2009). In particular, Triassic localities with abundant herbivore and carnivore coprolites are reported from United States (Case, 1922;Ash, 1978;Lucas et al., 1985;Nesbitt, 2001;Hunt and Lucas, 2005), Russia (Ochev, 1974), Poland (Bajdek et al., 2014;Zatoń et al., 2015), India (Matley, 1939a(Matley, , 1939bSohn and Chatterjee, 1979;Jain, 1983;Vijaya et al., 2009), Brazil (Souto, 2001;Dentzien-Dias et al., 2012;Da Silva et al., 2014) and Argentina (Rusconi, 1947(Rusconi, , 1949Mancuso et al., 2004;Hollocher et al., 2005;Fiorelli et al., 2013). ...
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... Hunt et al. (1994) stated that recurrent assemblages of coprolites are found in similar depositional settings and proposed the term 'coprofacies' for ichnofacies that are recognized on the basis of coprolites. This term was applied in several subsequent papers (Hunt et al., 1998(Hunt et al., , 2007Hunt and Lucas, 2005), but only a few researchers followed this idea. For vertebrate coprolites, it was applied by Hollocher et al. (2005) who cited Chin (1996, unpublished Ph.D. dissertation), and K. Chin is in fact the second author in Hunt et al. (1994). ...
... The Heteropolacopros ichnofacies is characterized by the presence of microspiral heteropolar coprolites that occur in fluvial redbeds. This ichnofacies occurs at least from the Early Permian (Hunt et al., 2005b, c) until the Late Triassic. ...
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The diverse ichnofaunas of the Late Triassic have been studied for almost 200 years. During the Late Triassic, facies favorable for the preservation of trace fossils were the result of low sea levels, monsoonal climates and the development of extensive depositional basins as Pangea began to fragment. The most abundant vertebrate trace fossils in the Late Triassic are tetrapod tracks, including Brachychirotherium, Chirotherium, “Parachirotherium,” Synaptichnium, Atreipus, Grallator, Eubrontes, Banisterobates, Trisauropodiscus, Evazoum, Tetrasauropus, Pseudotetrasauropus, Eosauropus, Apatopus, Batrachopus, Rhynchosauroides, Gwyneddichnium, Procolophonichnium, Chelonipus, Brasilichnium and Dicynodontipus. There are five tetrapod footprint biochrons of Triassic age that can be identified across the Pangaean footprint record. Coprolites are the second most abundant vertebrate trace fossils in the Late Triassic and include Heteropolacopros, Alococoprus, Dicynodontocopros, Liassocoprus, Saurocoprus, Strabelocoprus, Malericoprus, Falcatocoprus and Revueltobromus. Coprolites are useful in biochronology in the Late Triassic. Consumulites, dentalites (new term for bite marks), and burrows are moderately common in the Late Triassic. Nests and gastroliths are rare. All groups of vertebrate trace fossils demonstrate different diversity and abundance patterns through the Phanerozoic. Most vertebrate trace fossils have their earliest occurrences in the Devonian. The early Permian is an acme for both tracks and coprolites. The Late Triassic yields abundant tracks and coprolites, and tracks are also common in the Early Jurassic. The Jurassic and Cretaceous represent the times with the greatest diversity of vertebrate traces (tracks, coprolites, consumulites, dentalites, nests and gastroliths). The Quaternary also represents a time of vertebrate ichnological diversity (tracks, coprolites, regurgitalites, nests and burrows).
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